Coleoptera: Carabidae) from the Finisterre Range, Papua New Guinea

Mecyclothorax kavanaughi sp. n. (Coleoptera: Carabidae) from the Finisterre Range, Papua New Guinea

James K. Liebherr

Mecyclothorax kavanaughi sp. n. is described from the Finisterre Range, eastern Papua New Guinea. Mecyclothorax kavanaughi and M. toxopei Darlington synapomorphously exhibit setae on the fifth elytral interval, supporting their recognition as adelphotaxa. Mecyclothorax toxopei is known only from a unique male collected on Puncak Trikora, or Mt. Wilhelmina, Papua, Indonesia; a locality highly disjunct from the only known locality of M. kavanaughi. The precinctive distribution of M. kavanaughi supports recognition of the Finisterre Range as a distinct area of endemism, previously indicated by distributions of Cicadidae (Homoptera), Lycaenidae (Lepidoptera), and birds of paradise (Aves: Paradisaeidae). The geological accretion of the Finisterre terrane 3.0-3.7 Ma establishes a vicariance- based maximum age of origin for the clade including M. kavanaughi and M. toxopei. Prof. James K. Liebherr, Department of Entomology, Cornell University, Ithaca, NY 14853-2601, U.S.A. [email protected]

Introduction localities is consistent with our extremely incomplete Mecyclothorax Sharp, 1903 comprises species distrib- knowledge of the New Guinea fauna. In this paper I uted across Australia, New Guinea, Java, Borneo, describe a species discovered in the Finisterre Range, St. Paul’s and Amsterdam Islands, Lord Howe and Papua New Guinea by D. H. Kavanaugh, Califor- Norfolk Islands, New Caledonia, and the Society nia Academy of Sciences. This species’ adelphotaxon and Hawaiian Islands (Baehr 2008, Liebherr 2008). is distributed far away on Puncak Trikora, Papua, Though the Society Islands’ and Hawaiian Islands’ Indonesia. radiations are most diverse, with 67 species described Criteria supporting the recognition of the Finisterre from the former (Perrault 1992) and 125 from the Range as a distinct area of endemism are reviewed, latter (Britton 1948, Liebherr 2005, 2006, 2008), with the geological history of this portion of New the greatest morphological disparity and greatest Guinea used to establish an age of origin for the clade number of generalized species, characterized by mac- comprising the new species and its adelphotaxon, roptery, occur in Australia (Moore 1984). As pres- M. toxopei Darlington. ently constituted, Mecyclothorax contains the bulk of the species comprising the tribe Mecyclothora- cini; part of the subfamily Psydrinae as classified by Material and methods Moore (1963). Taxonomic material was obtained from the Califor- Knowledge of the New Guinea Mecyclothorax fauna nia Academy of Sciences (CASC). Resultant para- has been developed by sequential descriptions types are deposited in the CASC, Cornell University of species, often known from only single speci- Insect Collection (CUIC), and the National Museum mens, with each species known from only single or of Natural History, Naturalis (formerly Rijksmu- nearby localities (summarized in Baehr 2008). This seum van Natuurlijke Historie), Leiden (RMNH). highly allopatric, indeed disjunct array of species Diagnosis of the new species was facilitated by

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photographs of the unique male holotype of the closest putative relative, M. toxopei, held in the RMNH. Dissection and staining techniques follow Liebherr (2005, 2006, 2008). Male specimens were relaxed in near boiling water including a drop of Kodak Photo- flo detergent. The genital bulb was removed with fine forceps after cutting intersegmental membranes with sharpened minuten nadeln. The aedeagus and associated structures were cleared in cold 10% KOH overnight, neutralized in dilute acetic acid, and viewed in glycerine. Female reproductive tracts were dissected and cleared using like techniques, and then the reproductive tract and hindgut associated with the eighth abdominal segment were stained using Kodak Chlorazol Black dissolved in methyl cellos- olve. The stained dissections were cleaned of tracheae and mounted in glycerine on a microslide. Photographs of M. kavanaughi were taken using a Microptics Inc. photographic apparatus, using transmitted light for the genitalia, and transmitted plus direct light for the external views of the specimens. Stacked digital images were merged using Com- poseZ5 (Hadley 2006). Comparison photographs of the M. toxopei male holotype were provided by Dr. Jan van Tol (RMNH). Size is measured using standardized body length: the sum of the median length of the head from anterior margin of labrum to cervical ridge, plus median pronotal length, plus distance from the juncture of the raised scutellar apex with its flattened base to the apex of the longer elytron (if different bilaterally). Eye convexity is described using the ocular ratio; the Fig. 1. Mecyclothorax kavanaughi sp. n., / paratype maximum lateral distance between the outer surfaces (CUIC); scale bar, 1.0 mm. of the eyes over the minimum width of the frons between the eyes. Male genitalic terminology follows Liebherr (2006, 2008), whereas female genitalic terminology is based on Shpeley et al. (1985). #89-48 // D.H. Kavanaugh collector, CALIF. ACADEMY OF SCIENCES COLL. // PAPUA NEW GUINEA EXPEDITION – 1989 // HOLO- Taxonomy TYPE ? Mecyclothorax kavanaughi J.K. Liebherr Mecyclothorax Sharp 2008 (red bordered label) (CASC). Paratypes: 7?,5/. Same data as holotype (5? 4/ Mecyclothorax Sharp, 1903: 243 (type species Cyclothorax CASC, 1? 1/ CUIC, 1? RMNH). montivagus Blackburn by Andrewes 1939). A complete generic synonymy is provided in Liebherr Diagnosis (2008). Shiny, piceous beetles with convex, oviform elytra (Fig. 1), standardized body length 4.0-4.8 mm. This M. toxopei Mecyclothorax kavanaughi Liebherr, sp. n. species shares with Darlington, among all other known Mecyclothorax species, the presence of Figs 1, 3, 4–8 dorsal elytral setae in both the third and fifth elytral intervals. In addition, both species are characterized Type material. Holotype ?, Papua New Guinea, by a very short sinuation of the basolateral prono- Madang Province, Finisterre Range massif 7.1 air tal margin anterad the basal pronotal seta. The new km WSW of Teptep, 3450 m, 27 Mar. 1989, Stop species differs from M. toxopei in: 1, very reduced

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ot 3 aml mf 0,25 mm

Fig. 4. Mecyclothorax kavanaughi male aedeagus, right view: aml, aedeagal median lobe; dp, dorsal plate; lp, left paramere; mf, microtrichial field of internal sac; ot, ostial triangle; rp, right paramere.

Description Coloration. Dorsal body surface uniformly shiny piceous, with margins of elytral margins and epip- leura dark rufous; mandibular apices, maxillary and Figs 2-3. Elytral conformations of Mecyclothorax spp. labial palps, and antennomeres 1-3 smoky brunne- – 2, Scutellum and basal portions of M. toxopei median ous, apical antennomeres piceous; ventral surface elytral intervals illustrating isodiametric microsculp- with piceous basal coloration, legs and posterior ture, bilateral presence of parascutellar seta (pss, right margins of visible abdominal ventrites 3-6 paler, seta broken off), and shallow, smooth elytral striae; rufobrunneous. 3, M. kavanaughi scutellum and median elytral intervals Head. Vertex broadly convex, frontal grooves shal- illustrating reduced microsculpture, absence of paras- low but distinct, linearly convergent from anterior cutellar setae, and punctate elytral striae. Dorsal elytral supraorbital seta to fronto-clypeal suture which is setae (des) in third and fifth elytral intervals characterize obsolete between frontal grooves, frontal area laterad both species. frontal groove longitudinally convex, bordered laterally by deep clypeo-ocular prolongation (Larochelle & Larivière 2007); two supraorbital setae present microsculpture on the dorsal body surface, with each side; eyes small, moderately convex, ocular head, pronotum and elytral surfaces shiny (Figs. 1, lobe behind eyes broad and separated from gena by 3), not with evident microsculpture (Fig. 2); 2, pro- narrow but evident groove, ocular ratio 1.39-1.42; notal base covered with distinct punctures, each sep- antennae robust, pubescent from fourth antenno- arated from the adjacent punctures by shiny cuticle, mere to apex, antennomeres 7-10 about 1.5× long as versus pronotal base smooth with indistinct punc- broad; mentum tooth well developed, apex extended tures in deepest portions of laterobasal depressions in nearly as far anterad as lateral lobes, subacute, very M. toxopei; and 3, male aedeagal tip acutely angulate narrowly rounded. (Fig. 4) versus bluntly rounded in M. toxopei. Pronotum. Broad, lateral marginal depressions

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narrow, disc convex nearly to margin in apical ¾ Male genitalia. Aedeagal median lobe robust, broad, of length (Fig. 1); basolateral margin very briefly apex extended to an acuminate tip (Fig. 4); a small sinuate immediately anterad articulatory socket of ostial triangle (Liebherr 2006) present; aedeagal basal seta; base evenly convex between hind angles, internal sac with broad diffuse field of microspi- unmargined, covered with deep, distinct punctures cules covering euventral surface; dorsal plate large, that extend to very small, nearly smooth laterobasal its length about 0.5× distance from parameral apices depressions; median longitudinal impression consist- to aedeagal tip. Parameres diagnostic for Mecyclot- ing of linear row of punctures on pronotal base, obso- horacini (sensu Moore 1963), with right paramere lete though traceable on disc, terminated anteriorly broader, though elongate, and with setae lining ven- at very deep, smooth anterior transverse impressions tral surface, and left paramere attenuated apically, that define a strongly convex anterior callosity; front with only two short setae at apex. angles rounded, slightly extended anteriorly, lateral marginal depression widest just behind front angle. Female reproductive tract. Bursa copulatrix elon- Prosternal projection medially depressed between gate, about 4× as long as gonocoxal length, apically broad lateral margins that are continuous with mar- expanded (Fig. 5); spermatheca fusiform, spermath- ginal bead of procoxal cavity. ecal duct joined to bursa dorsad juncture of bursa Elytra. Disc convex, elevated relative to lateral and common oviduct. Gonocoxae with basal gono- margins; elytral striae 1-6 and 8 basally punctate coxite bearing three apical setae, two laterally and (Fig. 3), though very shallow to coplanar with adjoin- one medially, plus smaller microsetae along medial ing intervals between the large punctures, striae shal- margin, one at the apicomedial margin (Fig. 6); api- lower, smoother apically, obsolete at elytral apex; cal gonocoxite subtriangular, with two lateral ensi- seventh stria obsolete, evidenced basally by a series of form setae plus a dorsal ensiform seta, and two apical minute, very shallow punctures; parascutellar striole nematiform setae. indicated by series of five to six punctures in some individuals (e.g. Fig. 3), by a smooth impression in Distribution (Fig. 7) others; parascutellar seta absent in most individuals Known only from the type locality in the Finis- (present bilaterally in one, and present on left side terre Range, Huon Peninsula, eastern Papua New only in a second individual) (single known speci- Guinea. men of M. toxopei has parascutellar seta bilaterally present, Fig. 2); dorsal elytral setae present in both Habitat (Fig. 8) third and fifth intervals in nearly all individuals, D.H. Kavanaugh (pers. comm.): “The specimens from 3-5 (usually 4) setae in third interval, and from … were collected along a small, swift-flowing stream 0-3 setae in fifth interval (single specimen lacking at the bottom of a deep V-shaped valley above seta in fifth interval on left side has one seta present treeline. The valley itself was on a North-facing slope, on right side); eighth stria with series of lateral elytral with perhaps an artificially low treeline. Vegetation setae divided into two groups, an anterior series of on the slopes and ridges of the valley was low and 5-7 setae and a posterior series of 6 setae, posterior dense and included Spinifex and other grass-forms, seta in anterior series may be in a relatively isolated low shrubs, and scattered tree ferns. The Mecyclot- position between the two series; both a subapical and horax were found in gravel and under small rocks on apical seta present near elytral apex. flat banks about 0.5 m back from the water’s edge, Pterothorax. Mesepisternal concavity ventrad elytral in areas where tussocks of grasses were interspersed humerus lined with 10-12 minute, isolated punc- with open gravelly/rocky flats (hence relatively stable tures, intervening surface shiny, several more diffusely bars).” distributed punctures posteriorly near mesepimeron; metepisternum short, quadrate, medial and ante- Etymology rior margins of subequal length. Metathoracic flight The species epithet honors the collector, Dr. David wings vestigial, the flaplike vestigium about as wide H. Kavanaugh, who has greatly enhanced our as long, not extended beyond posterior margin of knowledge of the world’s montane and alpine cara- metathorax (two dissected females examined). bid beetle diversity. Legs. Tarsomeres short, moderately expanded apically (Fig. 1), the fourth metatarsomere with very short Identification apical lobes; tarsomeres 1-4 with inner and outer This species can be identified by modifying the most ventrolateral series of three to four long, stout setae, recent key to New Guinea Mecyclothorax (Baehr the setae slightly longer than depth of tarsomeres at 2008). Reiterating what Baehr noted for all other setal insertion. previously known species, this species is allopatric

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5 6

afs sg sp ams

les 0.25 mm gc1 0.10 mm des

gc2 ans

Figs 5-6. Mecyclothorax kavanaughi female reproductive tract and gonocoxae, ventral view. 5, bursa copulatrix and gonocoxae: bc, bursa copulatrix; co, common oviduct; gc1 and gc2, basal and apical gonocoxites; sg, spermathecal gland; sp, spermatheca. 6, female gonocoxa: afs, apical fringe setae of gonocoxite 1; ams, apicomedial seta of gc1; ans, apical nematiform setae of gonocoxite 2; des, dorsal ensiform seta; les, lateral ensiform setae.

Fig. 7. Distributional ranges of M. toxopei () and M. kavanaughi ().

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- Dorsal body surface shiny, microsculpture obsolete on dorsum of head, and on pronotal and elytral discs (Fig. 1); pronotal median base covered with distinct punctures; parascutellar seta absent in most individuals (Fig. 3). Finnistere Range, eastern Papua New Guinea (Fig. 7) . . . M. kavanaughi sp. n.

Discussion The hypothesized adelphotaxon relationship of M. kavanaughi and M. toxopei is based on shared possession of dorsal elytral setae in the fifth elytral interval. Such extraneous setae are not known from any other Mecyclothorax species. In addition, these two species exhibit three to five setae in the third elytral interval. Such increased setal numbers in the third interval, over the plesiomorphic presence of two dorsal elytral setae, are also observed in several other New Guinea species; M. kubor Baehr (2008) with five, M. julianae Baehr (1995) with four, and M. bilaianus Baehr (1998) and M. sedlaceki Darling- ton (Baehr 1995) with two or three setae present. However, the similarities in body size, pronotal shape and setation, and elytral setation shown between M. kavanaughi and M. toxopei all point to their closest phylogenetic affinity among described Fig. 8. Habitat surrounding alpine stream type local- species. ity of M. kavanaughi, Finisterre Range, Papua New The occurrence of M. kavanaughi in the Finisterre Guinea. Collecting locale was along stream just above Range supports recognition of this range as an area of white water rapids in center right of photo. Photograph endemism (Crisci et al. 2003). This mountain range courtesy David H. Kavanaugh. has been long recognized as housing precinctive species. Heads (2001) summarized the distributions of birds of paradise (Aves: Paradisaeidae), interpret- with all other New Guinea Mecyclothorax species. ing the disjunct occurrence of several species in the Insertion of the following couplets will provide trou- Finisterre Range and the Vogelkop region in a vicari- ble-free identification of the described taxon, but ance framework. Astrapia rothschildi Foerster of the the great geographic distance between the ranges of Finisterre Range is advanced as the adelphotaxon this species and its adelphotaxon M. toxopei, coupled to A. nigra (Gmelin) of the Vogelkop. Similarly, with the demonstrated flight-wing vestigialization of Paradisaea guilielmi Cabanis of the Finisterre is M. kavanaughi, strongly suggest that further species placed as sister species to the Vogelkop’s P. rubra of this clade remain to be discovered. Daudin. These latter two species, in turn, share 1. Elytral intervals 3 and 5 with setiferous punc- synapomorphous feather characters with P. decora tures (rarely individuals unilaterally lacking Salvin and Godman of the D’Entrecasteaux Islands. setae in fifth interval) ...... 1a Heads interprets these distributions and phyloge- - Only elytral stria 3 with setiferous punctures . . 2 netic relationships as the results of vicariance medi- 1a. Dorsal body surface with well-developed ated by lateral strike-slip movements of Melanesian microsculpture, the median elytral intervals terranes that have sequentially accreted onto the covered with isodiametric mesh (Fig. 2); northern margin of the New Guinea orogen. Among pronotal median base smooth, with indis- already docked terranes, that bearing the current tinct punctures present only laterally mesad Finnisterre Range is estimated to have accreted onto laterobasal depressions; parascutellar seta New Guinea most recently, about 3.0-3.7 Ma. present. Wilhelmina Top (Puncak Trikora), The distributions and phylogenetic relationships central Papua (former Irian Jaya), at 4200 m of cicadas have also been interpreted in such a (Fig. 7) ...... M. toxopei Darlington vicariance framework, with the Huon Peninsula or

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Finisterre Range being the possible source of the Guinea (Insecta, Coleoptera, Carabidae, Psydrinae). generic-level radiation classified as the genus Scotto- – Spixiana 21: 21-24. tympana Boer (Boer & Duffels 1996). Baehr, M., 2008. Two new species of the genus Mecyclotho- Among lycaenid butterflies, three species in the rax Sharp from New Guinea (Coleoptera: Carabi- Psychonotis caelius species group share similar pat- dae: Psydrinae). – Tijdschrift voor Entomologie 151: terns of distribution and endemism (Müller 2003); 133-140. P. hebes (Druce) distributed in the New Guinea cen- Boer, A.J. de & J.P. Duffels, 1996. Historical biogeography tral highlands, P. finisterre Müller in the Finisterre of the cicadas of Wallacea, New Guinea and the West Range, and P. marginalis Müller in the Owen Stanley Pacific: a geotectonic explanation. – Palaeogeography, Range. Palaeoclimatology, Palaeoecology 24: 153-177. Britton, E.B., 1948. A revision of the Hawaiian species of Though our knowledge of Mecyclothorax diversity Mecyclothorax (Coleoptera: Carabidae). – Occasional is no doubt in a very primitive state, the vicariance Papers of the Bernice P. Bishop Museum 19: 107- model incorporating accretion of Melanesian ter- 166. ranes onto New Guinea permits dating the common Crisci, J.V., L. Katinas & P. Posadas, 2003. Historical Bio- ancestor of M. toxopei and M. kavanaughi as no older geography. – Harvard University Press, Cambridge, than the time that the terrane bearing the currently 250 pp. uprising Finisterre Range sutured onto New Guinea: Hadley, A., 2006. CombineZ5 (computer program for 3.0-3.7 Ma (Heads 2001). Given only the currently stacking digital images). – Free Software Foundation described, putative sister species we cannot deter- Inc., Boston, MA www.hadleyweb.pwp.blueyonder. mine how these taxa relate to others in New Guinea. co.uk/CZ5/combineZ5.htm) [visited on: 4 February However as the center of diversity for the remain- 2008]. ing species is concentrated in the central highlands Heads, M., 2001. Birds of paradise, biogeography and (Baehr 2008)—including M. bilaianus, M. julianae, ecology in New Guinea: a review. – Journal of Bioge- and M. kubor, all exhibiting derived enhancement of ography 28: 893-925. Larochelle, A. & M.-C. Larivière, 2007. Carabidae (Insec- dorsal elytral setation—the outgroup to the M. toxo- ta: Coleoptera): synopsis of supraspecific taxa. –Fauna pei-M. kavanaughi clade is more likely to be distrib- of New Zealand 60: 1-188. uted there. Moreover, the likelihood that additional Liebherr, J.K., 2005. New species of Mecyclothorax (Cole- undiscovered species closely related to M. toxopei optera: Carabidae, Psydrini) from Polipoli, Maui de- and M. kavanaughi occupy the mountain ranges in fine an area of endemism on Haleakala Volcano, Ha- addition to the central highlands and the Finisterre waii. – Journal of the New York Entomological Society Range cannot be discounted. 113: 97-128. Liebherr, J.K., 2006. Taxonomic revision of the Mecy- clothorax beetles (Coleoptera: Carabidae, Psydrini) of Acknowledgements Molokai, Hawaii and recognition of areas of ende- David H. Kavanaugh kindly provided the detailed mism on Kamakou volcano. – Journal of the New York description and photograph of M. kavanaughi’s type Entomological Society 114: 179-281. locality. I thank Jan van Tol for photographs of the Liebherr, J.K., 2008. Taxonomic revision of Mecyclothorax unique holotype of M. toxopei Darlington, which Sharp (Coleoptera: Carabidae) of Hawaii Island: ram- pant genitalic evolution in a nascent island radiation. allow both diagnosis of the new species and better – Deutsche Entomologische Zeitschrift 55: 19-78. access to the diagnostic features of M. toxopei. Dan Moore, B.P., 1963. Studies on Australian Carabidae (Cole- Polhemus provided the New Guinea base map. This optera) 3. – The Psydrinae. – Transactions of the Royal research was supported in part by N.S.F. REVSYS Entomological Society, London 115: 277-290. grant DEB-0315504. Moore, B.P., 1984. Taxonomic notes on some Australasian Mecyclothorax Sharp (Coleoptera: Carabidae: Psydri- nae) and descriptions of new species. – Journal of the References Australian Entomological Society 23: 161-166. Andrewes, H.E., 1939. Papers on Oriental Carabidae. Müller, C.J., 2003. Three new species of Psychonotis Tox- – XXXV. On the types of some Indian genera. – An- opeus (Lepidoptera: Lycaenidae) from Papua New nals & Magazine of Natural History (11) 3: 128-139. Guinea. – Australian Entomologist 30: 159-165. Baehr, M., 1995. The genus Mecyclothorax Sharp, 1903 Perrault, G.G., 1992. Endemism and biogeography among in New Guinea (Coleoptera, Carabidae, Psydrinae). Tahitian Mecyclothorax species (Coleoptera: Carabidae: – Mitteilungen der Münchner entomologischen Ges- Psydrini). – In: G.R. Noonan, G.E. Ball & N.E. Stork ellschaft 85: 3-19. (Eds.), The biogeography of ground beetles of moun- Baehr, M., 1998. A further new species of the ge- tains and islands: 201-215. Intercept, Andover. nus Mecyclothorax Sharp from western New Sharp, D., 1903. Coleoptera, Caraboidea. – In: D. Sharp

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(Ed.). Fauna Hawaiiensis 3: 175-292. University Press, Cambridge. Shpeley, D., R.B. Madge, & G.E. Ball, 1985. The genus Celaenephes Schmidt-Göbel: characteristics, species, and relationships (Coleoptera: Carabidae: Lebiini). – Proceedings of the Entomological Society of Wash- ington 87: 283-307.

Received: 28 July 2008 Accepted: 31 Aug 2008

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